THE INFLUENCE of the DIET of the ARGENTINE STEM WEEVIL, Listronotus
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This thesis may be consulted by you, provided you comply with the provisions of the Act and the following conditions of use: you will use the copy only for the purposes of research or private study you will recognise the author's right to be identified as the author of the thesis and due acknowledgement will be made to the author where appropriate you will obtain the author's permission before publishing any material from the thesis. THE INFLUENCE OF THE DIET OF THE ARGENTINE STEM WEEVIL, Listronotus bonariensis (KUSCHEL) (COLEOPTERA: CURCULIONIDAE) ON THE FITNESS OF ITS P ARASITOID, Microctonus hyperodae LOAN (HYMENOPTERA: BRACONIDAE): PROSPECTS FOR 'INDIRECT' CONSERVATION BIOCONTROL I ._:-l_.;r__ ~'_.:... -:~ A thesis submitted in partial fulfilment .,-,---_ .. of the requirements for the Degree of Master of Applied Science at Lincoln University by Mauricio Alvaro Urrutia Correa Lincoln University 2005 Abstract of a thesis submitted in partial fulfilment of the requirements for the Degree of M. Appl. Sc. THE INFLUENCE OF THE DIET OF THE ARGENTINE STEM WEEVIL, Listronotus bonariensis (KUSCHEL) (COLEOPTERA: CURCULIONIDAE) ON THE FITNESS OF ITS P ARASITOID, Microctonus hyperodae LOAN (HYMENOPTERA: BRACONIDAE): PROSPECTS FOR 'INDIRECT' CONSERVATION BIOCONTROL By M. A. Urrutia Correa The Argentine stem weevil, Listronotus bonariensis (Kuschel) (Coleoptera: Curculionidae), is the most important pest species in New Zealand, costing the pastoral industry an estimated NZ$78-251 m per year (Prestidge et al., 1991). The parasitoid wasp Microctonus hyperodae Loan (Hymenoptera: Braconidae) was released in New Zealand in 1991 as a potential biological control agent of the weevil (Goldson et al., 1994). As M hyperodae is a pro ovigenic species, its lifetime complement of eggs is set at adult emergence (Phillips, 1998). Adult wasp fitness is likely to be affected by host nutrition, as resources carried over from immature stages to the adult stage of parasitoids can vary with the size and nutritional quality of the host (Jervis et al., 2001). The fecundity of Argentine stem weevil can increase after the addition of pollen to its diet (Evans. and Barratt, 1995). Conversely, a decrease in gonad development (i.e., vitellarium development and percentage of females with oocyte resorption) can occur after consumption of ryegrass containing endophyte (Barker and Addison, 1996). This research investigates tri-trophic-Ievel interactions on parasitoid fitness (i.e., egg load and body size) by assessing field-collected adult weevils, and by providing bee-collected pollen, buckwheat (Fagopyrum esculentum Moench cv. Katowase) pollen, ryegrass-only (Lolium multiflorum L. cv. Tama) , or ryegrass (Lolium multiflorum Lambrechtsen cv. Aries) containing endophyte (Neotyphodium lolii (Latch, Christensen and Samuels) Glenn, Bacon and Hanlin in the host diet in the laboratory. A survey was conducted in the Lincoln area to what extent pollen was part of the diet of field collected Argentine stem weevils. The frequency and types of pollen grains consumed were ii analysed. Also, gut fullness (in terms of ryegrass particle content), parasitoid larval presence, weevil gender and weevil gonad development were also assessed. Argentine stem weevil adults fed on pollen in the field, although a low proportion of them contained pollen in their gut. Gonad development was positively correlated with the extent of pollen consumption (i.e., number of individuals with sexually mature gonads). A cage experiment was conducted in the laboratory. Potentially parasitised adult Argentine stem weevils were collected from ryegrass on roadsides in the Lincoln area during 2004. They were randomly allocated to four treatments: ryegrass-only (control), ryegrass plus bee collected pollen, ryegrass plus buckwheat pollen, and ryegrass with endophyte, each replicated four times. Water was provided in every treatment. The parasitoids that emerged were assessed for size (i.e., hind leg tibial length) and potential fecundity (i.e., egg load) (see Phillips and Baird, 2001). Despite the fact that the addition of bee-collected or buckwheat pollen to the host diet had no significant effect on the number of adult parasitoids emerged, their body size or egg load (P > 0.05), some factors that could have affected a potentially positive correlation between parasitoid fitness and pollen consumption are discussed. Similarly, the addition of ryegrass with endophyte had no significant effect on the number of adult parasitoids emerged, their body size or egg load (P > 0.05). This leads to the suggestion that biological control (i.e., M hyperodae) and plant resistance (i.e., endophyte) strategies can work in a complementary way in integrated pest management programmes. The concept of benefiting the third trophic level with 'resource subsidies' made available selectively to the second level has the potential to lead to the suggestion of a new mechanism for enhancing parasitoid fitness, that is 'indirect' conservation biocontrol. Keywords: Argentine stem weevil, Listronotus bonariensis, Microctonus hyperodae, ryegrass, buckwheat, pollen, endophyte, conservation biological control, tri-trophic interactions. iii To my wife and daughters .. ......... , " ......... far from being a purely passive victim, obliterated without a trace, the host is often able to impress its mark ...... upon the insect parasitoid that destroys it" (Salt, 1941 ). iv Acknowledgements I would like to thank my wife Denise for her constant support and advice. Thanks as well to my daughters Valentina and Magdalena, who despite their young age allowed Daddy to read, write and think during those peak times. Thanks as well to the NZAID scholarship granted by the New Zealand Government for the opportunity to study this Master's Degree. Also I would like to thank my supervisor, Professor Steve Wratten, from Lincoln University for his unconditional professional and personal support. Thanks also to my co-supervisor, Craig Phillips, from AgResearch for sharing his expertise. Thanks as well to Bruce Chapman from Lincoln University, for his generous knowledge and important contribution in this Masters Degree. Many thanks also' to Mark Wade for his technical and intellectual contribution to this thesis. Also, thanks to David Baird, from AgResearch for his help in the statistical analysis of the data. Also thanks to Neville Moar from Manaaki Whenua Landcare Research for his help in the pollen identification. Finally, thanks to all the staff and students aligned to the National Centre for Advanced Bio-Protection Technologies who helped me in several aspects of the present work. v Contents Page Abstract / keywords 11 Acknowledgements v Contents VI Tables x Figures x Plates X111 Chapter 1: General Introduction 1 Biological control background 1 The Argentine stem weevil and current control practices background 3 Argentine stem weevil 3 Graminaceous crops and Argentine stem weevil in New Zealand 3 Management tools for Argentine stem weevil 6 Chemical control 6 Cultural control 6 Plant resistance 7 Biological control 8 Goal and Objectives 12 Hypotheses 12 Chapter 2: Gut contents of field-collected Argentine stem weevils 13 Introduction 13 Materials and Methods 14 1. Pollen content 14 2. Ryegrass particle content 15 vi 3. Gonad development in Argentine stem weevil 15 4. Parasitism 17 5. Statistical analyses 17 Results 17 Pollen content 17 Amount of pollen in the weevils' gut 17 Effects of pollen consumption on gonad development 19 Types of pollen 22 Ryegrass particle content 24 Parasitism 25 Effects of parasitism on weevils' gonad development and rye grass intake 25 Discussion 28 Pollen content 28 Argentine stem weevil diet breadth 28 Quantity of pollen in the weevils' gut 28 Effects of pollen consumption on gonad development 29 Types of pollen 32 Ryegrass particle content 34 Parasitism 35 Effects of parasitism on weevils' gonad development and ryegrass intake 36 Chapter 3: Effects of host diet on components of fitness of Microctonus hyperodae Loan 38 Introduction 38 Materials and Methods 40 Experiment 1 40 1. Experiment set up 40 vii 2. Rearing plants for Argentine stem weevil in experiment 1 40 3. Argentine stem weevil cages 40 4. Parasitoid morphological measurement 41 5. Quantifying egg load of the parasitoids 41 Experiment 2 43 6. Experiment set up 43 7. Methods during and after exposure to parasitoids 44 8. Rearing plants for Argentine stem weevil in experiment 2 45 9. Gonad development in the Argentine stem weevil 45 10. Gut fullness of the Argentine stem weevil 46 11. Exposure of weevils to adult parasitoids 47 12. Statistical analyses 47 Results 47 Experiment 1 47 Parasitoid tibia length and egg load 47 Experiment 2 50 Diet consumption by the weevil 50 Survival and fitness-related data of Argentine stem weevil 52 Components of fitness of the parasitoid 56 Number of adult parasitoids emerged 60 Discussion 60 Experiment 1 61 Parasitoid body size and egg load 61 Experiment 2 63 Diet consumption by the weevil 63 Survival and fitness of Argentine stem weevil 65 viii Fitness of the parasitoid 68 Number of adult parasitoids emerged 72 Chapter 4: General discussion 75 The host's mark 75 Implications on biological control 77 Selectivity issues 78 Is there room for improvement? 78 References 80 ix Tables Page Table 1. Combination of treatments of experiment 2 43 Table 2. The relationship between tibia length and egg load for parasitoids emerged from weevils fed with four treatments 58 Table 3.